Savoring The Past Positive Memories Evoke Value Representations in The Striatum

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Neuron. 2014 November 19; 84(4): 847856. doi:10.1016/j.neuron.2014.09.028.
Savoring the past: Positive memories evoke value

representations in the striatum
Megan E. Speer, Jamil P. Bhanji, and Mauricio R. Delgado
Department of Psychology, Rutgers University, Newark, NJ 07102, USA
Summary
Reminders of happy memories can bring back pleasant feelings tied to the original experience,
suggesting an intrinsic value in reminiscing about the positive past. However, the neural circuitry
underlying the rewarding aspects of autobiographical memory is poorly understood. Using fMRI,
we observed enhanced activity during the recall of positive relative to neutral autobiographical
memories in corticostriatal circuits that also responded to monetary rewards. Enhanced activity in
the striatum and medial prefrontal cortex was associated with increases in positive emotion during
recall and striatal engagement further correlated with individual measures of resiliency. Striatal
response to the recall of positive memories was greater in individuals whose mood improved after
the task. Notably, participants were willing to sacrifice more tangible monetary rewards in order to
reminisce about positive past experiences. Our findings suggest that recalling positive
autobiographical memories is intrinsically valuable, which may be adaptive for regulating positive
emotion and promoting better well-being.
Introduction
Sometimes it can be good to live in the past. Autobiographical memories serve a variety of
adaptive functions, such as bolstering a sense of self identity (Bluck et al., 2005;
DArgembeau and Van der Linden, 2005) or planning for the future (Schacter and Addis,
2007). When it comes to remembering positive life events, however, one adaptive function
of autobiographical memories may be its most prominent: it feels good to remember the
good times. That is, reminders of the past can bring back emotions tied to the original
experience (e.g., Bower, 1981; Westermann et al., 1996). For example, remembering a game
winning goal in a championship game may trigger a re-experience of positive emotions
associated with that day. In this way, the recall of positive autobiographical memories may
be intrinsically valuable to an individual by increasing or maintaining positive feelings that
contribute to ones general well-being.
2014 Elsevier Inc. All rights reserved.

Address correspondence to: Mauricio R. Delgado, PhD, Department of Psychology, 101 Warren Street, Smith Hall - Room 301,
Newark, NJ 07102, delgado@psychology.rutgers.edu, Phone: 973-353-3949, Fax: 973-353-1171.
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Speer et al.
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The capacity to savor or maintain positive feelings is associated with an increased ability for
emotion regulation (Carl et al., 2013), which in turn may promote successful adaptation to
stress (i.e., resilience; Philippe et al., 2009). Individuals afflicted with depression tend to
recall fewer autobiographical memories, especially positive ones (Young et al., 2013), and
have difficulties sustaining positive emotions (Carl et al., 2013; Heller et al., 2009).
Critically, these patients also show aberrant activation patterns in neural circuits involved in
reward processing (Delgado, 2007; Haber and Knutson, 2010; ODoherty, 2004), such as
reduced striatum responses to rewards (Pizzagalli et al., 2009) and positive feedback (Elliott
et al., 1998), as well as difficulty sustaining reward-related activity in the ventral striatum in
response to positive stimuli (Heller et al., 2009). Together, these observations highlight the
benefits of savoring events that elicit positive emotions to ones well-being, and underscore
the importance of understanding the neural mechanisms underlying changes in positive
emotion.
One potential way of eliciting positive emotions that engage reward-related circuitry may be
to recall pleasant autobiographical memories. Such memories tend to be vivid and rich (e.g.,
Schaefer and Philippot, 2005; Talarico et al., 2004) and benefit from having a positive value
attached to it during encoding (Scimeca and Badre, 2012) that is re-experienced during
recall (Westermann et al., 1996). In fact, a positive, arousing event (e.g., the potential to earn
money) can impact the encoding of memory in general (e.g., a neutral scene), engaging
activity in reward-related regions such as the midbrain and the ventral striatum at the time of
memory formation that promotes better subsequent recall (Adcock et al., 2006), potentially
via interactions with memory-related regions such as the hippocampus (Wittmann et al.,
2005). Thus, it is plausible that increased dopamine release during encoding influences
dopaminergic targets (such as the striatum) and deem the memories more relevant,
strengthening them over time and making them easier to access in the future (for review see
Shohamy and Adcock, 2010).
Autobiographical memories may also be adaptive in the sense that recalling the past is
something we already do naturally. Importantly, we may be able to capitalize on the positive
emotion we feel when reminiscing about happy memories, which unlike other forms of
emotion regulation (e.g., cognitive reappraisal) may be a proactive rather than reactive
strategy for increasing positive emotion. Indeed, the rewarding aspects of autobiographical
memories are a likely component of affective well-being and protection from disorder, but
their underlying neural substrates are poorly understood. That is, do autobiographical
memories display properties in common to other rewards? In particular, how does the brain
represent rewarding aspects of memories (i.e., positive valence, positive emotion during
recall)? Further, can autobiographical memories be assigned a value that is comparable with
other rewards (i.e., will people sacrifice a tangible reward for a more highly valued
autobiographical memory)?
In this experiment, we utilized functional magnetic resonance imaging (fMRI) to examine
neural activity during the recall of autobiographical memories, particularly memories with
positive content, which can lead to the re-experience of positive emotion. Participants first
participated in Day 1, where they were presented with common life event cues (e.g., Family
Vacation) and asked to describe specific memories of positive (e.g., visiting Disneyland) and
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neutral (e.g., packing for a trip) content, but not negative (e.g., lost luggage). Participants
who met criteria (i.e., had enough memories) were then invited to participate in Day 2 the
fMRI session. In the scanner, they recalled (i.e., brought the details of the original event to
mind) a number of positive and neutral memories prompted by the cues from the first
session, and provided ratings of valence (positive, neutral), feeling and intensity. We
hypothesized that recalling a positive memory would a) evoke positive feelings and b)
engage reward-related neural circuitry including the striatum, which is involved in the
processing of primary, secondary and abstract rewards (e.g., food, money and pleasant
music, Salimpoor et al., 2011, respectively; see for review Delgado, 2007; Haber and
Knutson, 2010; ODoherty, 2004) as well as active coping processes related to dealing with
negative affect (Delgado et al., 2009; Ledoux and Gorman, 2001). Finally, we conducted a
behavioral experiment on a separate cohort of participants to probe for the value of recalling
such positive memories. We hypothesized that participants would value the experience of
recalling positive autobiographical memories and consequently choose such an experience
over a more tangible reward (i.e., money).
Results
The recall of autobiographical memories elicits positive emotion

Nineteen healthy participants (10 females; mean age = 26.1, SD = 7.78) performed a cued
recall task while undergoing fMRI. Prior and following the task, participants were given the
Positive and Negative Affective Scale (PANAS; Watson et al., 1988) to assess mood change
during the experiment. They concluded the fMRI session by performing a card task for
monetary rewards (Delgado et al., 2000) which served as an independent localizer of
reward-related brain activity.
The autobiographical memory recall task involved the presentation of 21 positive and 21
neutral cues for a total of 42 trials. On each trial (Figure 1), participants were presented with
a written cue (e.g., Family Vacation) which prompted a memory a personal experience
from the past that was deemed as positive (e.g., visiting Disneyland) or neutral (e.g., packing
for a trip). The memories were previously validated in an interview three days earlier by
participants descriptions and ratings of emotional intensity and feeling (Day 1 see
procedures in Methods). Memory cues selected for the recall task were those in which the
valence rating (e.g., positive) matched their emotion ratings (e.g., high intensity, high
feeling). Upon seeing a cue (e.g., family vacation) participants had 14 seconds to recall a
memory. They made a button press to indicate the onset of the recall experience, and another
button press to indicate when they were done recalling the memory a procedure similar to
(Daselaar et al., 2008; Sharot et al., 2007). Participants were then asked to rate the memory
they just recalled with respect to valence (positive or neutral), level of emotion or feeling
(i.e., how did you feel when you recalled this memory; 1-4: 1 = neither good nor bad, 4 =
very good) and emotional intensity (i.e., how intense was the particular memory; 1-4: 1 =
not intense, 4 = very intense).
During performance in the memory cued recall task, we observed that positive memories
elicited an increase in experienced positive emotions. Recalled memories that were rated as
positive evoked greater emotional intensity (t(18) = 9.51, p < .001) and positive feeling
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(t(18) = 20.0, p < .001) compared to memories rated as neutral. Perhaps unsurprisingly,
participants spent more time during the recall of positive (M =6.7 s, SD = 3.0 s) compared to
neutral memories (M = 5.6 s, SD = 2.6 s; t(18) = -4.91, p < .001). There was no significant
relationship between feeling ratings and duration for positive (r(18) = -.18, p = .47) or
neutral memories (r(18) = .23, p = .34), however, suggesting that the observed enhanced
emotion cannot necessarily be due to a longer period of recall. Importantly, the onset of
memory recall did not differ between positive (M = 3.0 s, SD = 0.9 s) and neutral memories
(M = 2.9 s, SD = 1.0 s), suggesting that neither memory valence was easier or harder to
remember.
Neural activity represents the valence and emotion of autobiographical memory recall
We were interested in investigating the neural correlates underlying the recall of

autobiographical memories which elicited such subjective increases in positive emotion. We
performed 3 key analyses (see Methods). The first analysis was a random-effects whole
brain general linear model (GLM) and focused on the period of recall. We used the cluster
level statistical threshold plugin in BrainVoyager to correct for multiple comparisons
(Goebel et al., 2006). Resulting statistical maps were set to a threshold of p<0.005 and
corrected to a whole brain cluster correction threshold of p < .05 with a threshold of 6
contiguous voxels (162mm3 as determined by the plugin).
A simple contrast of positive compared to neutral autobiographical memories revealed
increased activity in corticostriatal circuits involved in reward-related processing (Balleine
et al., 2007; Delgado, 2007; Haber and Knutson, 2010; ODoherty, 2012). Specifically, this
analysis identified voxels within the striatum (including bilateral head of the caudate),
ventral medial prefrontal cortex, orbitofrontal cortex and anterior cingulate as showing
greater activity for positive compared to neutral autobiographical memories during time of
recall (see Table S1; Figure 2A-B; Figure S1). These ROIs were then used in a subsequent
follow-up analysis to test relationships between neural activity and emotion ratings collected
during the autobiographical memory task in the scanner.
Of particular interest was the involvement of the striatum and prefrontal cortex, given their
involvement in reward processing, in the recall of positive autobiographical memories. We
hypothesized that such regions would be modulated by the degree of positive emotion
evoked by the specific memory (i.e., feeling ratings on a trial by trial basis). We also
explored the possibility that these neural regions may be modulated by the degree of arousal
evoked by the memory (as indicated by emotional intensity ratings). To test this, we
conducted two additional tests (one for feeling ratings and one for emotional intensity
ratings during the memory task) on regions of interest (ROIs) in the striatum and prefrontal
cortex identified in the main contrast of positive and neutral memories. Specifically, we
included subjective ratings (of either feeling or emotional intensity) as a parametric
modulator for positive memory trials in a GLM and extracted parameter estimates from
these ROIs. This test was not biased by the ROI selection procedure because the parametric
modulator in each model was not correlated with the positive and neutral memory
regressors.
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We observed that positive emotion (i.e. feeling ratings) significantly modulated activity in
the caudate bilaterally (right: t(18) = 2.43 p = .03; left: t(18) = 2.38, p = .03; Figure 2C) and
medial prefrontal cortex (mPFC; t(18) = 2.40, p = .03; Figure 2D). This suggests that, during
the recall of positive life events, neural activity in these regions increases to the degree that
positive emotions were generated. In contrast, the second GLM with emotional intensity
ratings as a parametric regressor for positive memory trials did not significantly modulate
activity in these regions. These results suggest that activity in the striatum and mPFC during
the recall of positive autobiographical memories is associated with the magnitude of positive
emotions experienced as opposed to merely reflecting an arousal response.
Given this linear relationship between positive emotion and activity in regions previously
implicated in reward processing, one hypothesis is that the level of such engagement during
recall may be related to individual differences in a stress-protective factor, such as
resilience. That is, the ability to recall positive autobiographical memories in a manner that
increases positive emotion and recruits activity in these regions may benefit successful
adaptation from adversity. Highly resilient individuals tend to report greater levels of
subjective wellbeing, which may be the result of experiencing more positive emotion (e.g.
optimism) or engaging in active coping mechanisms (e.g. humor, mindfulness; Wu et al.,
2013) a process that has been associated with striatum function during aversive learning
(Delgado et al., 2009; Ledoux and Gorman, 2001). We observed that participants with
higher self-reported resiliency (Connor-Davidson Resiliency scale; Connor and Davidson,
2003) exhibited greater activity in the right caudate ROI defined by the contrast of positive
and neutral memory recall during the recall of positive autobiographical memories
(correlation between resilience scores and parameter estimates from the right caudate ROI:
r(19) = 0.458, p = .05).
Changes in mood after recalling autobiographical memories are related to striatal activity
In a second key analysis, we explored whether remembering positive memories would also
lead to mood improvements. We asked participants to report their current mood state using
the Positive and Negative Affective Scale (PANAS; Watson et al., 1988) before and after
the memory task. No significant changes in mood were observed in the sample (t(18) =
0.500, p = .62). However, there were individual differences as some participants moods
improved (n = 7), some worsened (n = 9) and some showed no change (n = 3). We then
conducted an exploratory analysis comparing behavioral and neural changes underlying the
mood-increase and the mooddecrease groups. Both groups were similar in terms of task
performance during recall, showing no differences in emotional ratings (positive memories:
t(14) = 0.798, p = .44; neutral memories: t(14) = 1.70, p = .11), or recall duration (positive
memories: t(14) = 0.076, p = .94; neutral memories: t(14) = 0.200, p = .84). Thus a question
remained about what was underlying the changes in mood across the experiment for these
individuals.
Based on a specific hypothesis that striatum activity underlies mood increases brought about
by positive autobiographical memory recall, we tested neural differences within this region
between individuals whose mood improved and individuals whose mood worsened after
recalling memories. To avoid potential biases within ROIs previously defined by the
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contrast of positive and neutral memories in our first analysis, we instead applied a small
volume correction (SVC) in a structurally defined striatum mask (including putamen,
caudate, and nucleus accumbens using the Harvard-Oxford probabilistic atlas; Desikan et
al., 2006), and then performed a Valence (positive/neutral) by Mood Change (moodincrease/mood-decrease) ANOVA. We observed a significant interaction in the right ventral
striatum (x, y, z: 10, 11, 1; F(1,14) = 17.55, p < .05, SVC; Figure 3), driven by a marked
increase in activity for positive relative to neutral memories in the mood-increase group (t(6)
= 11.46, p < .001), but no significant change for the mood-decrease group (t(6) = 0.32, p = .
76). We also observed a significant interaction in the left putamen (x, y, z: -25, 1, -3; F(1,14)
= 18.64, p < .05, SVC), with the mood-increase group showing a similar increase for
positive compared to neutral memories (t(8) = 4.29, p = .005), and the mood-decrease group
trending in the opposite direction (t(8) = 1.75, p = .12). Although the results should be
deemed exploratory, given the small sample size of the groups, these relationships suggest
that, in certain individuals, an increase in striatum activity during the recall of positive life
events can accompany an improvement in mood.
Ventral striatum responses distinguish between the valence of monetary outcomes and
memory recall
The third key analysis performed addressed an interesting question of whether brain regions
that actually distinguish between more tangible rewards (i.e., money) during reward
paradigms would also distinguish between positive and neutral autobiographical memories
during recall. We asked participants to play a surprise card game for monetary rewards
(adapted from Delgado et al., 2000) at the conclusion of the memory task. Consistent with
previous findings using this reward paradigm (see for review, Delgado, 2007), a contrast of
gain and loss outcomes revealed robust activation in the striatum (Figure 4; Table S2). We
defined this reward-based ROI in the right ventral striatum and extracted parameter
estimates for positive and neutral memories. An increase in activity related to the recall of
positive compared to neutral autobiographical memories was observed in the right ventral
striatum (t(18) = 2.33, p = .03). Thus, voxels in a ventral striatum ROI functionally defined
by a contrast between monetary gains and losses (i.e. tangible, extrinsic reward) were also
sensitive to the valence of autobiographical memories in the same individuals.
Positive autobiographical memories are valuable

Our findings suggest that recalling positive experiences from the past increases ones
positive emotion and engages reward-related neural circuitry, such as the striatum and
medial prefrontal cortex. One implication of these results is that reminiscing about positive
experiences is intrinsically valuable to an individual. If this is the case, however, then people
should be willing to forgo another, more tangible reward (i.e., money) for the opportunity to
recall positive as compared to neutral autobiographical memories. We tested this hypothesis
in a behavioral experiment using a separate cohort of participants.
The behavioral study adopted the same design as the fMRI study with one important
difference: prior to recalling a memory, participants had a choice of which memory to
conjure up. They chose between a positive (e.g., Family Vacation) or a neutral (Grocery
Shopping) memory cue (Figure 5A). Both options were accompanied by a specified
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monetary payoff that ranged from $0.01 to $0.04 (every combination of monetary payoffs
was represented in option pairs as in similar paradigms; Hayden et al., 2007; Tamir and
Mitchell, 2012). Thus, some choices required participants to choose to forego a larger
monetary payoff in order to recall their preferred memory in the task. Participants then
recalled the selected memory as in the fMRI study, and at the end of the task received a
monetary payoff according to the sum of their choices.
Twenty-six healthy participants (19 females; mean age 20.0, SD = 2.98) preferred to recall
positive over neutral memories. Compared to chance (50%), specifically, participants chose
positive memories 85% of the time when choices were of equal value (t(25) = 10.8, p < .
001) and 70.7% of the time across all relative payoffs (t(25) = 7.45, p < .001). To measure
the subjective value associated with choosing a positive over a neutral memory, we
calculated the point of subjective equity (PSE) for each participant by fitting a cumulative
normal distribution to their choice behavior (adapted from Tamir and Mitchell, 2012; Figure
5B). The PSE represents the monetary payoff foregone to choose to recall a preferred
memory. That is, PSE is the difference in monetary payoffs between the two options when a
participant was equally likely to choose a positive or neutral memory (i.e. where the
cumulative normal distribution passed 50% and limited to the range of relative payoffs: $0.03 to +$0.03). Participants were willing to forgo an average of 1.94 cents (SD = 1.22) to
recall a positive instead of a neutral memory, (t(25) = -8.16, p < .001), which was still
significant when controlling for gender (t(25) = 2.66, p = .014). That is, participants
sacrificed 28% of potential monetary earnings to recall positive rather than neutral
memories.
Given the known deficits in reward-related processing and positive emotion observed in
depressed individuals (Carl et al., 2013; Pizzagalli et al., 2009), we also tested whether
participants willingness to forgo monetary rewards to recall positive memories was related
to symptoms of depression. Interestingly, there was a significant positive correlation
between PSE and depression scores (Beck Depression Inventory; Beck et al., 1961; r(25) = .
39, p = .047, and when controlling for gender, partial r(25) = .40, p = .048). Individuals with
greater symptoms of depression were less willing to give up money to recall a positive
experience.
Discussion
Autobiographical memory is remarkable not only for the ability to re-experience the past,
but also for allowing us to rekindle emotions captured during the original experience. Our
findings suggest that reminiscing about positive past experiences produces internally
generated positive emotions, which may be rewarding in and of itself. In the current study,
the recall of positive autobiographical memories elicited an increase in positive emotion, as
measured by subjective ratings of feelings, and recruited neural circuits involved in rewardrelated processing. Activity in the striatum and medial prefrontal cortex was modulated by
the strength of positive emotion experienced and striatal activity in particular correlated with
individual measures of resiliency. Further, re-living these happy memories in ones mind led
some participants to improve their mood during the experiment, with such individuals also
showing greater ventral striatum activity when recalling positive compared to neutral
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autobiographical memories. Finally, a separate cohort of participants chose to recall positive

compared to neutral memories even if it came at a financial cost; that is, they were willing to
choose recalling a positive memory over a neutral one even when the neutral memory was
accompanied by a monetary reward. This behavior suggests that individuals value positive
memories to the point of sacrificing more tangible rewards. Taken together, the findings
suggest that positive autobiographical memories can be intrinsically valuable to an
individual, serving adaptive functions such as bolstering positive emotion which is
significant for an individuals well-being and ability to cope with negative affect (Carl et al.,
2013).
The recruitment of reward-related regions such as the striatum and medial prefrontal cortex
during the recall of positive autobiographical memories and association with increased selfreported positive emotion is in line with our hypothesis that such memories are valuable and
elicit positive emotions attached to the original experience. Specifically, the striatum has
been involved in processing the subjective value of typical (Delgado, 2007; Haber and
Knutson, 2010; ODoherty, 2004) and atypical (Delgado et al., 2009; Ledoux and Gorman,
2001; Salimpoor et al., 2011; Tamir and Mitchell, 2012) rewards, and decreases in selfreported positive emotion correlate with decreases in striatal signal in depressed individuals
(Heller et al., 2009).
There are alternative interpretations of striatum recruitment during memory recall worth
considering, however. For instance, it is possible that striatal activity solely reflects
emotional arousal, which is a component of an emotional response. While it is conceivable
that positive emotional responses will elicit a higher arousal than neutral feelings, arousal
per se did not explain the modulation of the striatum activation. In particular, this region was
modulated by subjective ratings of emotional feelings, but not emotional intensity. It is also
possible that the striatum activity is due to re-assigning value to a recalled stimulus,
consistent with findings of striatum activity in post-choice changes in preference (Sharot et
al., 2009), when an increase in perceived value of a stimulus is observed after it is chosen.
This interpretation is not inconsistent with our account that such memories carry value and
serve to increase positive emotion and perhaps improve mood in select individuals.
The prefrontal cortex has been more consistently observed among studies of
autobiographical memory retrieval (Markowitsch et al., 2003; Piefke, 2003), but it is also
plausible that striatum activity is related to aspects of the memory process itself, such as
adaptive updating or the cognitive control of episodic retrieval (Scimeca and Badre, 2012),
rather than the account proposed in the current paper. Our design attempted to control for
aspects of the memory process itself by asking participants to recall, describe, and
emotionally rate all memories, both neutral and positive, three days prior to scanning.
Accordingly, there was no difference in the amount of time it took to bring a positive or a
neutral memory to mind in the subsequent scanning session, suggesting similar ease of
recall, and thus similar retrieval success, between positive and neutral memories. Further,
the current findings show a specific role for striatal activity in representing affective aspects
of memory and lesser engagement when recalling neutral memories. Although it is worthy
to note that participants may have re-encoded the memories with additional value during the
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initial retrieval process three days prior to the scanning, consistent with Scimeca and
Bradres framework of adaptive encoding (2012).
One interesting aspect of the observed striatal activity was the consistent observation of
greater responses during the recall of positive compared to neutral autobiographical
memories regardless of the location of the ROI within the striatum. The whole-brain contrast
of positive and neutral memories elicited activation in the caudate nucleus, part of the dorsal
striatum and involved in affective and cognitive aspects of behavior (Balleine et al., 2007;
Delgado, 2007; Haber and Knutson, 2010). However, we also observed greater responses to
the recall of positive memories in the ventral striatum. Specifically, we observed an
interaction between memory valence and mood increase in a structurally defined striatum
ROI during memory task performance. We also observed greater activation to positive
compared to neutral memory recall in a functionally defined independent ROI in the ventral
striatum related to reward processing (i.e., showing responses to monetary gains compared
to losses). These results highlight the involvement of both dorsal and ventral striatum in the
recall of positive autobiographical memories.

There are several important implications of our findings. First, we reinforce the idea that
positive autobiographical memories have an adaptive function to increase positive emotion,
which can potentially benefit ones ability to use emotion regulation (Carl et al., 2013).
Consistent with the idea that positive autobiographical memories carry intrinsic value, we
demonstrated that participants had a significant preference for recalling past, positive events,
at the point of forgoing a more tangible reward. This is somewhat surprising considering that
participants have access to their own positive memories at any time, and therefore are not at
a loss if they waive an opportunity to recall a positive memory in order to earn a larger
monetary reward. However, it fits with the idea that people tend to value experiences over
material possessions (e.g., Dunn et al., 2011), and that looking at the past positively, or more
nostalgically, can make people happier (Zhang and Howell, 2011). This may be a unique
feature of increasing positive emotions via autobiographical memories instead of other
methods. For instance, participants who are asked to create a positive mental image (e.g., a
baby laughing) have enhanced positive emotions much like those elicited by positive
memories, but are less willing to sacrifice money to imagine positive situations in
comparison to participants who forgo rewards to recall the positive past (see Supplemental
Information; Figure S2). Although this is just one example, and more research is necessary
to investigate other ways in which positive emotions can be elicited, these results support an
inherent value attached to the recall of autobiographical memories.
Second, we highlight a neural mechanism involved in increases in positive emotion and
mood related to recalling positive autobiographical memories a potentially important
emotion regulation strategy to cope with negative affect (Rusting and Dehart, 2000). Not
surprisingly, patients with depression have difficulties recalling positive autobiographical
memories (Young et al., 2013) and using this type of strategy (Dalgleish et al., 2013), which
may be related to an inability to sustain activity in the striatum during positive emotion
(Heller et al., 2009). We observed individual differences in mood improvement after
recalling positive memories, Although this finding is exploratory given the small samples, it
lends support to the idea that some people are able to benefit more than others when
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recalling the positive past, in accordance with individual differences in emotion regulation
success (e.g., Martin and Delgado, 2011). Interestingly, we also observed that resilience was
associated with greater striatal activity during the recollection of positive memories,
consistent with the thought that cultivation of positive emotions builds resources to enhance
life satisfaction, for example, by increasing the likelihood of experiencing positive emotion
in the future and even garnering ones resilience against negative affect (Tugade and
Fredrickson, 2004). Savoring positive events from our past may be an adaptive tool for
combating lifes troubles. Therefore, it is intriguing that resilient individuals, who have a
greater tendency to experience positive emotions, are also those who exhibit more rewardrelated activity in the striatum when remembering happy moments. In a similar vein,
individuals who showed a boost in mood from before to after the memory task had more
striatal activation when recalling positive memories. Although we cannot test a causal
relationship, this raises the question of whether purposefully engaging in positive
recollection of the past increases ones resilience, and perhaps mood, over time.
We also observed a pattern linked to depression in our behavioral study. Individuals with
greater depressive symptoms were less willing to give up money to recall a positive event.
Depression has been linked to not only functional anatomical deficits in autobiographical
recall (Young et al., 2013), but also a diminished ability to recall specific, positive events
(Williams and Scott, 1988). One explanation is a lack of normative positivity bias in
depression rather than a negativity bias, which our results seem to support (Suslow et al.,
2001). Therefore, it is possible that if recalling pleasant memories feels effortful and evokes
only a small, fleeting positive feeling, then depressed individuals may feel like the payoff is
too small. Our results suggest that reminiscing about past positive experiences may have an
adaptive role in regulating and maintaining positive emotion, which may promote better
well-being, particularly in the case of affective disorders like depression.
While experiencing positive emotion when reminiscing about happy memories is not an
entirely new idea, the uniqueness in regulating emotions using autobiographical recall is, in
part, that memories are an abundant resource we already have available to us, whereas other
emotion regulation strategies are typically stimulus-driven. For instance, studies examining
the cognitive regulation strategy of reappraisal (e.g., Wager et al., 2008) require the
reinterpretation of something negative to something less negative or positive. In contrast,
recalling the past is a natural phenomenon that people choose to do quite frequently in their
daily lives (Killingsworth and Gilbert, 2010) and therefore does not require instruction,
reinterpretation, or necessarily a cue. In this way, positive memories may uniquely represent
a proactive, rather than reactive method of generating positive emotion. Our findings
demonstrate not only that something naturally-occurring like recalling the past can
successfully influence mood in some individuals, but it is linked with activity in rewardrelated circuitry which fails to be sustained in depressed patients during experience of
positive emotions (Heller et al., 2009) and leads people to forgo an extrinsic reward
(money) in order to experience it.
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Methods
Participants
Twenty-eight healthy right-handed individuals were recruited via flyers posted on the
Rutgers campus to participate in a 2-day study (13 males, 15 females). Given our
experimental requirements (e.g., meeting criteria for number of available memories), we had
a number of exclusions prior to scanning. Specifically, participants completed an
autobiographical memory questionnaire on Day 1 and returned for the fMRI session on Day
2 only if they met criteria for inclusion (see procedures for Day 1; 23 met criteria). Five
participants did not make it to the fMRI session on Day 2 based on these criteria.
Additionally, we excluded 2 participants for excessive head motion (more than 4mm in any
direction), 1 for performance (i.e., 46% missed responses), and 1 withdrew before
completing the scanning session (claustrophobia). Final analysis was therefore performed on
19 participants (9 males, 10 females; mean age = 26.1, SD = 7.78). Our slightly higher
attrition rate is due to the potential for participants to be excluded after Day 1 as well as Day
2. All participants were free of neurological or psychological conditions, and were not
taking any medications at the time of testing. Participants gave informed consent in
accordance with the Rutgers University Institutional Review Board for the Protection of
Human Subjects in Research.
Experimental design
Day 1: Autobiographical Memory Questionnaire
Participants wrote about real memories prompted by each of 70 common life event cues
(e.g., Family Vacation). The written cues were a compilation of cues created for this study
and cues used in prior studies (Markowitsch et al., 2003; Piefke, 2003; Sharot et al., 2007).
For each cue, participants selected a memory in which they had been personally involved
and had occurred at a specific place and time. For each memory, participants reported a brief
description, the location, the date, and gave subjective ratings for valence (positive or
neutral), emotional intensity (i.e., how intense was the particular memory; 1-4: 1 = not
intense, 4 = very intense), and feeling (i.e., how did you feel when you recalled this
memory; 1-4: 1 = neither good nor bad, 4 = very good). Importantly, participants were
instructed to select memories that were positive (e.g. visiting Disneyland) or neutral (e.g.
packing for a trip), but not negative memories (e.g., lost luggage).
To be invited back to the imaging session (Day 2), individuals had to report at least 21
positive and 21 neutral memories. For each participant who met criteria, the 21 most
positive memories (i.e. high intensity, high feeling) and the 21 most neutral memories (i.e.
low intensity, low feeling) were selected as unique event cues to recall in the scanner.
Additionally, participants filled out the following questionnaires related to resiliency and
emotion: the Connor-Davidson Resiliency scale (CD-RISC; Connor and Davidson, 2003)
and the Beck Depression Inventory (BDI; Beck et al., 1961).
Speer et al.
Page 12
Day 2: fMRI session
Participants returned for the second session (Day 2) three days later. While in the MRI
scanner, participants first reported their current mood state via the PANAS (Positive and
Negative Affective Schedule; Watson et al., 1988), then completed a cued recall
autobiographical memory paradigm.
In the memory task, participants re-accessed 21 positive and 21 neutral memories triggered
by event cues from their questionnaire on Day 1. Each trial (Figure 1) included one written
event cue displayed for 14s. Participants were asked to recall the same memory from Day 1
and to elaborate on it for as long as they could or until the 14s were up. Participants made a
button press to indicate the start of a memory (i.e. when it began to form in their mind) and
another button press to indicate the end of a memory (if they finished elaborating on the
memory before time was up). After a delay of 2-4s, participants rated the memory on
valence, emotional intensity, and feeling (3s for each rating). An 8-10s delay separated one
trial from the next. The task was administered over the course of three functional runs, each
containing 7 positive and 7 neutral event cues. Cues were pseudorandomly presented such
that no more than 4 consecutive events of the same valence were displayed in a row.
Immediately afterwards, participants again rated their current mood state via the PANAS.
The pre- and post-task ratings of mood allowed for an assessment of mood change.
Specifically, mood change was calculated as the change in positive affect subscale (10
adjectives) from pre-autobiographical memory task to post- autobiographical memory task.
While still in the scanner, participants performed a surprise monetary reward task (the cardguessing game adapted from Delgado et al. (2000). This task occurred without warning as to
not influence the memory task and particularly the mood ratings post-memory task. The
purpose of this paradigm was to identify reward-related regions of interest (ROIs) to serve
as independent ROIs to test with positive and neutral memory regressors. In each trial of the
card task, participants saw a card with a question mark inside for 2s. They guessed whether
the cards value was higher or lower than the number 5 by making a button press. After a
short 2-4s delay, the card and monetary outcome were displayed. A correct response earned
a green checkmark signifying a gain of $1.00 whereas an incorrect response earned a red X
signifying a loss of $0.50. Unbeknownst to participants, outcomes were predetermined to
control schedule of reinforcement and number of gain and loss trials (20 each for a total of
40 trials).
At the conclusion of the scanning session participants were debriefed and compensated for
their time in the scanner and bonus money earned in the card game.
fMRI Data AcquisitionA 3T Siemens Magnetom Trio scanner was used for acquisition
of T1-weighted MPRAGE structural images (256 x 256 matrix, FOV = 256 mm, 176 1-mm
sagittal slices). Additionally, functional images were acquired in 35 contiguous obliqueaxial slices (3 x 3 x 3 mm voxels) prescribed parallel to the AC-PC plane with a single shot
gradient echo EPI sequence (TR = 2 s, TE = 25 ms, FOV = 192, flip angle 90, bandwidth =
2232 Hz/Px, echo spacing = 0.51) in three runs of the memory task and one run of the
monetary reward task.
Speer et al.
Page 13
Data were preprocessed and analyzed using BrainVoyager QX (v2.3, Brain Innovation).
Functional images were motion-corrected (six parameters), slice-timing corrected using a
cubic spline interpolation, and spatially smoothed using a Gaussian kernel of 8 mm FWHM.
Further, the data were temporally smoothed with voxelwise linear detrending and high-pass
filtering of frequencies (three cycles per time course). The images were spatially normalized
to the Talairach stereotaxic space (Talairach and Tournoux, 1988) in BrainVoyager.
fMRI Data AnalysisFunctional data were analyzed using a whole brain random-effects
general linear model (GLM). The memory task was modeled using two valence regressors
representing positive and neutral trials during memory recall and a regressor representing
missed trials (i.e., no valence rating given for the memory, 2.1% missed trials). The memory
regressors begin at memory formation and end after elaboration, with this period defined by
participants own button presses in each trial (for onset and conclusion of memory recall).
The monetary reward task was modeled using two regressors representing gain and loss
trials during the 2s outcome phase along with a regressor representing missed trials (no
response, 3.0%). For both the memory and monetary reward tasks, regressors were
convolved with a canonical double-gamma hemodynamic response function and six
regressors for motion parameters were included in the model.
To correct for multiple comparisons, we used the cluster level statistical threshold plugin in
Brain Voyager (Goebel et al., 2006) which uses Monte Carlo simulations in order to
determine the likelihood of observing clusters of various sizes in a given map at a set
threshold. After correction, the map automatically applies the minimum cluster size
threshold that produces the desired cluster-level false-positive alpha rate (5% was chosen).
We applied a voxel cluster threshold of 6 contiguous voxels (162mm3 as determined by the
plugin) defined at a threshold of p<0.005 to obtain a corrected alpha < 0.05. We then
performed 3 main analyses with the neuroimaging data.
1) Whole brain contrast of positive > neutral memories: We conducted a whole brain
contrast between positive and neutral memories during memory recall to examine the effect
of memory valence. From this simple contrast, we identified ROIs in regions previously
implicated in reward processing, namely the right caudate nucleus in the striatum and the
mPFC (for review see Haber and Knutson, 2010), and conducted follow-up analyses. First,
we tested whether positive emotion (i.e. feeling ratings or emotional intensity ratings)
modulated neural activity in the striatum during the recall of positive memories on a trial by
trial basis. Specifically, we conducted two separate analyses that included subjective ratings
(of either feeling or of emotional intensity during the recall task) as a parametric modulator
for positive memory trials in the GLM (orthogonalized with respect to the original positive
memory regressor). To test the relation of positive emotion to neural activity during recall, a
t-test was conducted on striatum and prefrontal cortex ROI parameter estimates for the
subjective feeling parametric modulation and the subjective emotional intensity parametric
modulation. Second, we examined individual differences in neural activity during the recall
of positive memories. In particular, we extracted parameter estimates from the striatum and
prefrontal cortex ROIs defined by the whole brain contrast of positive > neutral memories
Speer et al.
Page 14
and tested for correlation with self-reported scores of resiliency as measured with the
Connor-Davidson Resiliency scale (CD-RISC; Connor and Davidson, 2003).
2) Mood improvement and memory valence ANOVA using striatum mask: We probed
the relationship between neural activity during memory recall and mood change from
beginning to the end of the autobiographical memory task (i.e. post-PANAS pre-PANAS).
Participants were grouped into mood-increase (n = 7) and mood-decrease (n = 9) groups,
while participants with no mood change were excluded (n = 3). Based on a specific
hypothesis that striatum activity underlies mood increases brought about by positive
autobiographical memory recall we performed a Valence (positive/neutral) by Mood Change
(mood-increase/mood-decrease) ANOVA within a striatum ROI. We used a non-biased
structurally defined striatum mask (including putamen, caudate and nucleus accumbens
using the Harvard-Oxford probabilistic atlas; Desikan et al., 2006) and applied a small
volume correction (p < 0.05, corrected).
3) Monetary reward task contrast of Gains > Losses: We conducted a contrast of gain
and loss outcomes in the monetary reward task to identify reward-related ROIs (see Table
S2). The goal of this analysis was to confirm that a reward-related functionally defined
ROI would show an independent effect of memory valence. We used the functionally
defined reward ROIs in the striatum based on the contrast of gain > loss and ran a GLM
using positive and neutral memory recall regressors to examine overlap in neural activity for
positive autobiographical memory and a more tangible reward (i.e., money).
Behavioral Study
Participants
Thirty-two healthy individuals were recruited via the Rutgers psychology student subject
pool to participate in a 2-day study (9 males, 23 females). Similar to the fMRI study,
participants completed an autobiographical memory questionnaire on Day 1 and returned for
Day 2 only if they met criteria for inclusion (see procedures for Day 1; 26 met criteria). The
final sample included twenty-six participants (7 males, 19 females; mean age 20.0, SD =
2.98). Participants gave informed consent in accordance with the Rutgers University
Institutional Review Board for the Protection of Human Subjects in Research.
Experimental design day 1: Autobiographical memory questionnaire

The first session followed the same procedure as the fMRI study except the autobiographical
memory questionnaire included 84 (rather than 70) life events. Participants returned for Day
2 only if they described at least 35 positive and 35 neutral memories on their questionnaire.
For each participant who met criteria, the 35 most positive (i.e. high intensity, high feeling)
and 35 most neutral (i.e. low intensity, low feeling) memories were used in the memory
choice task on Day 2.
Experimental design day 2: Memory choice task
Participants returned three days later to perform the autobiographical memory choice task.
On each of 35 trials, participants made a choice between two options presented
Speer et al.
Page 15
simultaneously: 1) recalling a positive memory or 2) recalling a neutral memory; both

options were accompanied by a specified monetary payoff. The payoffs ranged from $0.01
to $0.04 and were presented in increments of $0.01 such that every combination of monetary
payoffs was represented in option pairs (as in previous studies Hayden et al., 2007; Tamir
and Mitchell, 2012). Thus, a participants choice of an option was based on a combination of
preference for recalling the cued memory as well as preference for receiving the specified
monetary payoff (in comparison to preference for the cued memory and monetary payoff in
the other option). Similar to the fMRI study, participants then recalled the selected memory
for 14s making button presses to indicate recall duration, followed by ratings of valence,
emotional intensity, and feeling.
Behavioral Data Analysis
To measure the subjective value associated with choosing a positive over a neutral memory,
we calculated the point of subjective equity (PSE) for each participant by fitting a
cumulative normal distribution to their choice behavior (adapted from Deaner et al., 2005;
Tamir and Mitchell, 2012). We first calculated the percentage of time participants chose a
positive over a neutral memory for each of the monetary differences between those two
options (i.e. relative payoffs of -$0.03, -$0.02, -$0.01, $0, +$0.01, +$0.02, and +$0.03). For
each participant, a cumulative normal curve was fit to the resulting seven data points using
the Matlab curve fitting toolbox. The PSE was calculated as the point where the cumulative
normal distribution passed 50%. This represents the relative monetary value (gain or loss)
when a participant was equally likely to choose a positive or neutral memory. PSE
calculations were limited to the range of relative payoffs (i.e. -$0.03 to +$0.03).
Supplementary Material
Refer to Web version on PubMed Central for supplementary material.
Acknowledgments
This research was supported by funding from the National Institute on Drug Abuse to M.R.D (DA027764) and a
National Science Foundation SBE Postdoctoral Research Fellowship to J.P.B. (1305994). We would like to thank
Elizabeth Tricomi, Susan Ravizza and Dominic Fareri for helpful comments and discussion, and three anonymous
reviewers for the constructive suggestions.
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Figure 1.
fMRI autobiographical memory task and behavior. (A) On each trial, participants recalled
either a positive or neutral memory while an event cue was displayed for 14s. Participants
indicated the beginning and end of the memory with button presses during the recall period.
Then, they gave subjective ratings of the memorys valence (i.e., positive or neutral),
emotional intensity, and feeling. (B) Positive memories were rated as significantly greater in
positive feeling and emotional intensity than neutral memories. P < 0.05, two-tailed test.
Error bars represent 1 s.e.m.

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Figure 2.
Neural activity during the recall of autobiographical memory. ROIs defined by contrasting
positive and neutral memory recall revealed activity in (A) the striatum, including the head
of the caudate, and (B) the medial prefrontal cortex. P < 0.05; corrected. (C) BOLD signal in
the right caudate (peak at x, y, z: 11, 1, 6) and (D) medial prefrontal cortex (peak at x, y, z:
-13, 58, 24) during the recall of positive versus neutral memory was parametrically
modulated by positive feeling ratings on each trial. The plots show BOLD signal parameter
estimates for positive memory recall according to the subjective ratings of feeling and
emotional intensity. Error bars represent 1 s.e.m. See also Figure S1, Table S1.

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Figure 3.
Ventral striatum activation during memory recall and subsequent mood change. (A) A
valence (positive/neutral) by mood change group (increase/decrease) ANOVA revealed a
significant interaction in the right ventral striatum. P < 0.05; SVC. (B) Mean parameter
estimates from the right ventral striatum are displayed showing greater activity for positive
compared to neutral memory in the mood-increase group (n = 7), but no significant change
in the mooddecrease group (n = 9). P < 0.05, two-tailed test. Error bars represent 1 s.e.m.

Speer et al.
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Figure 4.
Voxels within the ventral striatum that respond to the receipt of monetary reward also
distinguish between the valence of autobiographical memories during recall. (A) Monetary
reward ROIs defined by contrasting gain and loss outcomes in a card guessing task revealed
activity in the ventral striatum bilaterally. P < 0.05; corrected. (B) A monetary reward ROI
(i.e. the right ventral striatum) showed significantly greater activity for the recall of positive
memory as compared to neutral memory. P < 0.05, two-tailed test. Error bars represent 1
s.e.m. See also Table S2.

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Figure 5.
Memory choice task and behavior. (A) On each trial, participants made a choice between
two life event cues (positive, neutral) to recall, where each choice was associated with a
monetary payoff ranging from $0.01 to $0.04. The rest of a trial was the same as the fMRI
memory task. Participants recalled a memory while viewing the (selected) event cue for 14s,
made button presses to indicate the memorys beginning and end, and then gave subjective
ratings for the memory. (B) The graph represents the monetary value of positive
autobiographical memory for all participants in the behavioral study. Each dot represents the
percentage of trials where participants chose to recall a positive over a neutral memory (yaxis) for each relative payoff (x-axis). Relative payoffs were the difference between the
monetary value of the two options on a particular trial (i.e. positive neutral). We measured
the relative monetary value of recalling positive autobiographical memory by finding the
point where the cumulative normal distribution, fit to the data, passed 50% (Deaner et al.,
2005; Tamir and Mitchell, 2012). Across the sample, participants were willing to forgo 1.94
cents (SD = 1.22) to recall a positive over a neutral memory. See also Figure S2.


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